Trabajo Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) Nomenclature and taxonomic status of the listed by Philippi (1860) (: , Teiidae and Tropiduridae)

Jaime Troncoso-Palacios1, Yery Marambio-Alfaro2,3 1 Programa de Fisiología y Biofísica, Facultad de Medicina, Universidad de Chile, Independencia 1027, San- tiago, Chile. 2 Laboratorio de Sedimentología y Paleoambientes. Instituto de Ciencias Naturales Alexander von Humboldt, Facultad de Ciencias del Mar y Recursos Biológicos, Universidad de Antofagasta. Avda. Universidad de An- tofagasta 02800, Chile. 3 Parménides Limitada, Avda. Batallones de Atacama 112, Caldera, Atacama, Chile.

Recibido: 29 Diciembre 2019 ABSTRACT Revisado: 23 Marzo 2020 In 1860 Rudolph Amandus [Rodolfo Amando] Philippi published the book “Reise durch die Aceptado: 05 Mayo 2020 Wueste Atacama auf Befehl der chilenischen Regierung im Sommer 1853–54”, a wonderful work Editor Asociado: A. S. Quinteros of natural history, which was for a long time the most comprehensive source of geographic, cultural, botanic and zoological information on the Atacama Desert of Chile (currently Atacama

and Antofagasta regions). In this book, Philippi listed several species of lizards, and described doi: 10.31017/CdH.2020.(2020-060) five new species. However, the taxonomic identities of several of these species still remain to be clarified. Here we provide comments on these species and show that not all of Philippi`s (1860) type specimens were collected in Atacama Desert as has long been thought. We desig- nate lectotypes for Proctotretus pallidus and P. melanopleurus; and confirm that P. modestus is a junior synonym of L. bellii and that P. pallidus is a junior synonym of L. nigromaculatus. In the case of P. marmoratus, since it is a secondary homonym of L. marmoratus Gravenhorst, 1838, we consider it an invalid name.

Key Words: Aporomera, Atacama, Desert, Helocephalus, , Proctotretus.

RESUMEN En 1860 Rodolfo Amando [Rudolph Amandus] Philippi publicó el libro (sic) “Viage al Desierto de Atacama hecho de orden del Gobierno de Chile en el verano 1853-54”, un maravilloso tra- bajo de historia natural, el cual fue por mucho tiempo la fuente más completa de información geográfica, cultural, botánica y zoológica del desierto de Atacama de Chile (actualmente las regiones de Atacama y Antofagasta). En este libro, Philippi listo varias especies de lagartos, y describe cinco nuevas especies. Sin embargo, las identidades taxonómicas de varias de estas especies permanecen sin clarificar. Aquí nosotros proveemos comentarios sobre estas especies y demostramos que no todos los especímenes tipo de Philippi (1860) fueron colectados en el desierto de Atacama como se ha pensado por largo tiempo. Nosotros designamos lectotipos para Proctotretus pallidus y P. melanopleurus; y confirmamos que P. modestus es un sinónimo menor de L. bellii y que P. pallidus es un sinónimo menor de L. nigromaculatus. En el caso de P. marmoratus, dado que es un homónimo secundario de L. marmoratus Gravenhorst, 1838, lo consideramos un nombre inválido.

Palabras claves: Aporomera, Atacama, Desert, Helocephalus, Liolaemus, Proctotretus.

Introduction The German naturalist and explorer Rudolph [Ru- Atacama auf Befehl der chilenischen Regierung im dolfo] Amando Philippi (1808–1904), is well known Sommer 1853–54”, describing his journey and field for his major early contributions to the natural his- survey through the Atacama Desert carried out on tory of Chile. An overview of his life and work was behalf of the Chilean government between Novem- summarized by Kabat and Coan (2017). In 1860 ber 1853 and February 1855. The book provides data Philippi published the book “Reise durch die Wueste on various subjects as geography, culture, botany and Author for correspondence: [email protected] 175 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860) zoology with a detailed account of the trajectory. group”), which include the nigromaculatus, platei, Philippi`s journey in the Atacama Desert was wide- and other groups; and Eulaemus (or “Argentinean ranging and led from the coast to Andean highlands, group”), which include the darwinii, montanus, and covering the current Atacama and Antofagasta other groups (see Lobo et al., 2010). Also, Philippi regions of northern Chile (Fig. 1). In the Zoology (1860) included species currently assigned to the section of the book (Chapter 8, pp. 156–190), one genus Callopistes, which includes only two species of amphibian and eight species are discussed (pp. larger teiid lizards (Harvey et al., 2012), and from the 165–169), including the description of a new genus to genus Microlophus, which includes approximately and five new species providing a drawing of three of 20 species (Benavides et al., 2007) with several them (reproduced here in Fig. 2). Most of these liz- taxonomic problems among the Chilean species (Troncoso-Palacios, 2018). Almost all species listed by Philippi (1860) are controversial in regard to their taxonomic status or their occurrence in the Atacama Desert (Ortiz and Núñez, 1986). Hereafter, we review the lizards listed by Philippi (1860) on the basis of extant type specimens and comparative material from several Chilean collections and provide com- ments on their nomenclatural and taxonomic status.

Materials and methods

We examined three type specimens collected by Philippi and an additional seven type specimens of his collection were examined through photographs: Helocephalus nigriceps ZSM 38/1930 lectotype, Proctotretus modestus ZMB 5350, 70546–47, P. mel a - nopleurus FMNH 9969 syntype, and the specimens NMW 18914:1,2 considered by some authors as syntypes of Proctotretus pallidus (Tiedemann and Häupl, 1980; Gemel et al., 2019). For the specimens NMW 18914:1,2 we estimated our measurements based on the rule included in each photograph (in millimeters). We also reviewed photographs of six non type specimens collected by Philippi. For com- parisons, we examined 227 specimens not collected by Philippi but belonging to species that have been Figure 1. Sketch of the route followed by Philippi during his recorded in the area visited by him or in the sur- travel through the Atacama Desert. He started in Coquimbo (1, in November 1853) and traveled by ship to Caldera (3, in May roundings (Appendix I). Scales were observed using 1854), without visit Huasco (2), although some type specimens binocular lenses 0.8–5x. The characters for scalation has been claim as collected in this last locality (see comments were taken according to Pincheira-Donoso and in regards to Proctotretus pallidus). Then, he went to Copiapó (4, in May), returned to Caldera (3, in May). Later he went to Núñez (2005) and Troncoso-Palacios et al. (2015). Chañaral (5, in December), Taltal (6, in December) and Paposo (7, in December), sailed up to Mejillones (8, in December), Results and Conclusion returned to Taltal (6, in January 1855) and traveled to Tilopozo (9, in January) and San Pedro (10, in January). He returned, passing again through Tilopozo (9, in February), Pajonal (11, Proctotretus marmoratus (Philippi 1860, p. 165) in February), Finca de Chañaral (12, in February), and Puquios The genusProctotretus is currently considered a ju- (13, in February) before finally reaching Copiapó (4, in Febru- nior synonym of Stenocercus (Frost, 1992), a genus ary) at the end of his journey. that do not occur in Chile. However, currently all ards currently belong to the genus Liolaemus, which Chilean species formerly listed in the genus Procto- has been split into two subgenera, each split into tretus are placed in the genus Liolaemus (Etheridge, several groups: Liolaemus (sensu stricto or “Chilean 1995; Abdala and Quinteros, 2014), among them,

176 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021)

Figure 2. Lizards illustrated in Philippi (1860), here modified from two different copies of the book. Lateral aspect (right), cephalic scalation (center) and dorsal scalation (left). Above=Helocephalus nigriceps. Middle= Proctotretus bisignatus, listed as P. nigromaculatus in the zoological section of Philippi (1860). Below= P. pallidus.

P. marmoratus was considered as belonging to Philippi (1860) did not provide additional features Liolaemus by Boulenger (1885, p. 140) who suggest (e.g. color pattern or localities), no conclusion on that it is a questionable junior synonym of L. nitidus the species identity can be reached. (Wiegmann, 1834), an opinion shared by Ortiz and Núñez (1986) and Pincheira-Donoso and Núñez Proctotretus nigromaculatus (Wiegmann, 1834) (2005). Moreover, Philippi (1860) pointed out that (Philippi 1860, p. 166, tab. VI, Fig. 2; reproduced in P. marmoratus “the ventral part from the chin to here in Fig. 2) the anus has rounded and smooth scales”, the smooth Philippi (1860) starts his brief characterization of scales on the throat being a diagnostic trait of Lio- this species (currently Liolaemus nigromaculatus) laemus (Etheridge, 1995). Thus, the correct name attributing the species authority to Wiegmann, but combination is Liolaemus marmoratus (Philippi, then he added the following sentence in regards 1860), however, it is a secondary homonym with L. to the species illustration: “S(ection) Zool(ogy) marmoratus Gravenhorst, 1838 (ICZN, 1999, Art. tab(ularum) nomine Proct(otretus). bisignatus”, 53.3) and then Philippi`s marmoratus is an invalid which introduced a new name (P. bisignatus) as ju- (but available) name. We do not recommend the nior synonym of P. nigromaculatus. The reasons why proposing of a replacement name because the species he introduced this new synonym are unknown. As a identity cannot be established. Philippi (1860) did possible hypothesis to explain why Philippi listed this not provide a type locality for “P. marmoratus”, type taxon as P. nigromaculatus but provided a drawing of specimens are currently lost (Ortiz and Núñez, 1986) it with the legend P. bisignatus, we propose that per- and description is brief and inaccurate, making it haps Philippi intended to describe this species as “P. impossible to reach a clear conclusion on the species bisignatus” and made a plate before he realized that it identity. Boulenger (1885) may have proposed the had already been described as P. nigromaculatus by questionable synonymy of P. marmoratus and L. ni- Wiegmann. Later, Boulenger (1885, p. 147) synon- tidus because of Philippi’s (1860) statement "(dorsal ymized P. bisignatus with L. nigromaculatus. Müller scales are) very pointed and strongly keeled" in P. and Hellmich (1933a) restricted the type locality of marmoratus, as is found on L. nitidus, a diagnostic L. nigromaculatus to Huasco (Atacama region, Chile) feature for this last species (Pincheira-Donoso and and Müller and Hellmich (1933b) resurrected P. Núñez, 2005). Assuming that Philippi collected the bisignatus as a of L. nigromaculatus, res- specimens of “P. marmoratus” in his Atacama jour- tricting its type locality to Caldera (Atacama region, ney, we note that two species found in these localities Chile). Afterwards, several authors considered both, have strongly keeled dorsal scales: L. nitidus and L. nigromaculatus and L. bisignatus as full species L. zapallarensis Müller and Hellmich, 1933. Since (Ortiz, 1981; Pincheira-Donoso and Núñez, 2005),

177 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860) but Troncoso-Palacios and Garín (2013) corrected south of the Atacama Desert. Boulenger (1885, p. the type locality of L. nigromaculatus to the transect 141) included to P. modestus as a questionable ju- between Puerto Viejo and Copiapó and determined nior synonym of L. chiliensis, whereas according to that the population from Caldera (L. bisignatus) is Müller and Hellmich (1933a) and Hellmich (1934), conspecific withL . nigromaculatus. Despite the lack P. modestus is “identical” to L. altissimus altissimus of a description and being introduced as junior sy- Müller and Hellmich, 1932, this latter considered nonym, P. bisignatus (=L. bisignatus) is an available to be a junior synonym of L. bellii Gray, 1845, by name because a name associated with a drawing Núñez (2004). The synonymies of Boulenger (1885), and published before 1931 is available (ICZN, 1999, Müller and Hellmich (1933a), and Hellmich (1934) Art. 12.2.7) and a name introduced as synonym but were rejected by Ortiz and Núñez (1986) because L. used as available name before 1961 is also available chiliensis and L. altissimus altissimus do not occur in (ICZN, 1999, Art. 11.6.1). However, given that it is the Atacama Desert. In fact, there are no Liolaemus based in the same species that L. nigromaculatus, L. species from the mountains of near Santiago that also bisignatus (Fig. 3) is a junior synonym of L. nigro- occur in the localities visited by Philippi (Valladares- maculatus. A more detailed analysis can be read in Faúndez, 2011; Troncoso-Palacios, 2014), but Ortiz Troncoso-Palacios and Garín (2013). and Núñez (1986) were unaware that the types of P. modestus were not collected in the Atacama Desert Proctotretus modestus (Philippi 1860, p. 166) but rather near Santiago. Apparently, Philippi saw Philippi (1860) described it from both the Atacama a Liolaemus species in the Atacama Desert that he desert and the mountains of the Santiago Provin- erroneously confused with a L. bellii from the moun- ce. The syntypes were assumed lost by Ortiz and tains near Santiago, then he collected the specimens Núñez (1986), but without making reference to in this last locality and provide the description in his the syntypes mentioned by Müller and Hellmich Atacama book (Philippi, 1860), causing confusion (1933a). In fact, Müller and Hellmich (1933a) and and leading to Ortiz and Núñez (1986) to assume Hellmich (1934) listed three syntypes, two females that P. modestus occurs in the Atacama Desert. and one juvenile male, placed in the Zoologisches Since Müller and Hellmich (1933a) refer to Museum Berlin (currently Museum für Naturkunde ZMB 5350 as “type”, it should be considered as Berlin) with the number ZMB 5350 (currently ZMB lectotype and ZMB 70546–47 as paralectotypes. 5350, 70546–47) and they designated the current Our examination of these specimens (Fig. 4) allow ZMB 5350 as “type” because it is the larger one. It us to agree with Müller and Hellmich (1933a) and is currently thought that P. modestus was collected Hellmich (1934) in that it is conspecific with L. by Philippi in his journey by the Atacama Desert bellii, because P. modestus has lanceolate and su- (Ortiz and Núñez, 1986), however, the three type bimbricate dorsal scales, dorsal pattern formed by specimens were collected by Philippi in “Santiago “V” dark stripes bordered of whitish scales on the de Chile”, a locality from central Chile, far to the juvenile specimen, dorsal pattern formed by diffu-

Figure 3. Holotype of P. bisignatus (MNHNCL 1477) collected by Philippi and probably used to write the characterization of P. nigromaculatus and for drawing P. bisignatus.

178 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) se dark stripes bordered of whitish scales on adult 1860” was a secondary homonym (and then not a specimens and ventral dark reticulation in both valid name) with Liolaemus (Sauridis) modestus, juvenile and adults, all diagnostic characters of L. Tschudi, 1845. However, this last is a Stenocercus bellii (Pincheira-Donoso and Núñez, 2005). species (Laurent, 1984), then this claim is incorrect, Finally, we remark that when Müller and but both L. altissimus altissimus and P. modestus are Hellmich (1933a) and Hellmich (1934) addressed junior synonyms of L. bellii. the status of P. modestus, they decided to keep L. altissimus altissimus as the valid name, because they Proctotretus melanopleurus (Philippi 1860, p. 166) erroneously thought that “P. modestus Philippi, Philippi (1860) provided neither type locality, nor

Figure 4. Specimens of Proctotretus modestus. A), B) Lectotype ZMB 5350, adult female. C), D) Paralectotype ZMB 70546, adult female. E), F) Paralectotype ZMB 70547, juvenile male. All photographs by Frank Tillack.

179 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860) list of the type specimens nor an illustration of his Pincheira-Donoso and Núñez (2005, p. 455) provide P. melanopleurus. The description is brief (ten lines). different voucher number for the syntypes ofL . me- Boulenger (1885, p. 155), included P. melanopleurus lanopleurus (MNHNCL 1646, two specimens), in re- as questionable junior synonym of L. darwinii (Bell, gards to the numbers MNHNCL 1549–50 provided 1843), a proposal followed by Quijada (1916), who by Ortiz and Núñez (1986). Later, Pincheira-Donoso listed three syntypes placed in the Museo de Historia and Núñez (2007), remarked again on the doubtful Natural de Chile (MNHNCL) and he assumed the status of this species: “is an enigmatic taxon. Indeed, type locality as “Atacama”. Ortiz and Núñez (1986) it not only is known on the basis of two specimens, pointed out that two syntypes of P. melanopleu- but the type locality also still remains a mystery”. rus are placed in the Museo Nacional de Historia Pincheira-Donoso and Núñez (2007) also pointed Natural de Chile (MNHNCL 1549 and 1550) and out that the relationships between L. isabelae Na- the third in the Field Museum of Chicago (FMNH varro and Núñez, 1993, and L. melanopleurus are a 9969). They rejected Boulenger`s (1885) synonymy “matter for discussion”, suggesting the possible status because L. darwinii occurs in southern South Ame- of L. isabelae as junior synonym. Troncoso-Palacios rica in localities not visited by Philippi (1860) in his (2014) provided a list of lizards of Atacama Region Atacama travels through northern Chile. Although and stated that he reviewed one Liolaemus speci- Ortiz and Núñez (1986) pointed out a resemblance men (MZUC 11770, L. juanortizi Young-Downey to L. platei, they stated that they “prefer to consider and Moreno, 1992) from Puquios, Atacama, the its status (L. melanopleurus) as doubtful because last locality visited by Philippi before his return to the specimens are poorly preserved, which do not Copiapó, and since L. juanortizi has a dark lateral allow the observation of other features (besides the stripe as is found on L. melanopleurus, Troncoso- black lateral stripe and midbody scale counts) and Palacios (2014) suggested that MZUC 11770 may because no more specimens are known”. In regards be conspecific with L. melanopleurus. However, to the third specimen placed in the Field Museum of Puquios, which ranges from 1400 to 2000 meters Chicago, they pointed out that it is poorly preserved above sea level (masl), is out of the distributional and discolored, questioning its assignment to L. and elevation ranges of L. juanortizi, which has its melanopleurus. Later, Ortiz (1994) pointed out that closest record in Quebrada Patón, approximately 50 L. melanopleurus is an “enigmatic Philippi`s species”. km S from Puquios (Troncoso-Palacios, 2013) and Etheridge (1995) included L. melanopleurus in the it is only known from 3800 masl (Pincheira-Donoso L. montanus group, however, this was rejected by et al., 2008). We conclude that the information on Núñez et al. (2000) since tibia and supralabial scales the MZUC 11770 label is inaccurate and this speci- do no match with the features found in the L. mon- men of L. juanortizi was probably collected along a tanus group; they also stated that “this species (L. road that goes through Puquios to reach 3800 masl melanopleurus), if it exists, belongs to the chiliensis in some locality off the path followed by Philippi. group” (probably in reference to subgenus Liolae- Moreover, we did not find L. juanortizi in two field mus also known as the Chilean group). Pincheira- campaigns to Puquios (March 2016 and January Donoso and Núñez (2005, p. 333) stated that “the 2019), where we found only L. velosoi Ortiz, 1987. taxonomic position of L. melanopleurus is very Troncoso-Palacios et al. (2016) pointed out that “L. confusing, especially due to age of the three known melanopleurus remains a problematic species in specimens…, and their natural deterioration”, but terms of identification as the type locality is impre- proposed the “resurrection” of L. melanopleurus (p. cise and no additional specimens have been found 8). However, doubts arise on the application of the in more than 100 years”. Finally, Núñez and Gálvez term “species resurrection” by Pincheira-Donoso (2015) listed MNHNCL 1646, which include two and Núñez (2005) because, in , this term specimens, as the “holotype”, but without an indica- is applied when a taxon considered as junior synon- tion of which one of these two specimens they refer. ym is revalidated (e.g. Szederjesi et al., 2018), but We also remark that it is not possible to designate L. melanopleurus was not a junior synonym at the a holotype for L. melanopleurus, instead a lectotype time of Pincheira-Donoso and Núñez (2005). They should be designed. also stated that L. melanopleurus is not a synonym Philippi (1860) stated that P. melanopleurus of L. platei but that could be a senior synonym of L. has the following measures (in inches and lines maldonadae Núñez et al. 1991 (p. 443). Remarkably, converted here to cm): total length= 11.4 cm, head

180 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) length= 1.2 cm length, foreleg length= 1.7 cm, hind lack almost all the tail, the head of the lectotype is leg length= 2.3 cm, tail length= 6.8 cm, snout-foreleg slightly crushed (but cephalic scalation is discerni- distance= 1.7 cm, foreleg-hind leg distance= 2.1 ble). In regards to the FMNH 9969 specimen: SVL= cm. When we reviewed the syntypes placed in the 44.8 mm (Ortiz and Núñez, 1986); tail is complete; MNHNCL (May 2011 and November 2016), these no lateral dark stripe is discernible, and color pattern shared the number MNHNCL 1646, without dis- is severely deteriorated. tinction between specimens, however, both are easily We reviewed specimens of almost all Liolaemus distinguishable because one is much larger (snout (sensu stricto) that occur in the Atacama Desert vent length= 70.6 mm) than other (SVL= 46.7 mm). (except L. puna) and agree with Pincheira-Donoso Two different voucher codes should be provide for and Núñez (2005) in that it is unlikely that L. melano- the two specimens that currently shared the MN- pleurus could be assignable to L. platei or any of the HNCL 1646 voucher code, however, such action is species of the L. platei group, because in our review under the sole authority of the MNHNCL collection of 71 specimens of this group —L. nigrocoeruleus manager and beyond the scope of our study. The tail Marambio-Alfaro and Troncoso-Palacios, 2014, L. is currently lacking in both specimens; thus, total platei, L. velosoi and L. hellmichi Donoso-Barros, length and tail length cannot be compared with the 1975— we found a maximum SVL of 61.2 mm; data provided by Philippi (1860). Other measures whereas the lectotype of L. melanopleurus has SVL= are (MNHNCL 1646, larger and smaller specimen 70.6 mm. Moreover, we did not found a specimens respectively): head length= 1.6 and 1.2 cm, foreleg of the L. platei group with four scales on the frontal length= 2.2 and 1.6 cm, hind leg length= 3.3 and 2.4 area (2 anterior and 2 posterior) as occurs in the cm, axilla-ingle distance (foreleg-hind leg distan- lectotype of L. melanopleurus. On the other hand, ce?)= 2.6 and 1.9 cm. We have no data for FMNH L. paulinae Donoso-Barros, 1961, and L. puna Lobo 9969. This strongly suggests that Philippi (1860) took and Espinoza, 2004, appear to be unlikely candidates the measures from the smaller MNHNCL 1646 spe- because L. paulinae has maximum SVL= 54.6 mm cimen. The squamation described by Philippi (1860) (Pincheira-Donoso and Núñez, 2005) and L. puna is useless to differentiate between the type specimens has maximum SVL= 55.6 mm (Lobo and Espinoza, placed in the MNHNCL because both shared the 2004). Furthermore, L. nitidus can be discarded as features provided in the description (keeled and conspecific with L. melanopleurus because it has ovate shaped dorsal scales). However, only the larger 28–38 midbody scales, whereas L. melanopleurus specimen MNHNCL 1646 (Fig. 5) displays the deep has 42–56. black lateral stripes described by Philippi (1860), a If Philippi collected L. melanopleurus in the character that was used by him to name the species Atacama Desert as was assumed by Quijada (1916), (melanopleurus means black side in Greek). All this then the most probably candidates to be conspecific evidence suggests that Philippi based his description with it due to its size and midbody scale counts are of P. melanopleurus on data from at least two spe- L. atacamensis Müller and Hellmich, 1933 (max. cimens. Since the larger MNHNCL 1646 specimen SVL= 67.2 mm, midbody scales= 47–54), L. cons- features the black lateral stripe running from the tanzae Donoso-Barros, 1961, (max. SVL= 75.3 mm, shoulder to the middle of the trunk, we designate midbody scales= 54–64), L. isabelae (max. SVL= it as the lectotype. MNHNCL 1646 specimens have 82.8 mm, midbody scales= 54–60), L. nigromacula- the following features (lectotype and paralectotype, tus (max. SVL= 83.0 mm, midbody scales= 49–62) respectively): midbody scales 56, 42; 7, 6 supralabial and L. zapallarensis (max. SVL= 85.3 mm, midbody scales with the fourth turned upward; nasal and scales= 48–54); all of which have a dark lateral stripe rostral scales not in contact in the lectotype and in on males with additional color features that could contact in the paralectotype. The lectotype has four be assumes to be absent in L. melanopleurus type scales on the frontal area, 2 anterior and 2 posterior, specimens due the deteriorated conservation status whereas it is a single scale in the paralectotype (MN- of these, but L. zapallarensis can be discarded be- HNCL 1646) and we have no data for the FMNH cause the dorsal scales of it have keels and mucrons 9969 specimen. The MNHNCL 1646 paralectotype markedly more developed than the dorsal scales of has a short dark brown spot over the shoulder until L. melanopleurus. the axilla. Whitish belly in both specimens, without While it has been long assumed that L. mela- additional discernible color features. Both specimens nopleurus and all of Philippi`s other species were

181 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860)

Figure 5. Type specimens of Liolaemus melanopleurus. A), B), C) and D) MNHNCL 1646 lectotype. E) MNHNCL 1646 paralectotype. F) FMNH 9969 paralectotype (Photograph by Kathleen Kelly). collected during his Atacama journey, here we clarify identify “P. melanopleurus” despite one synonymy that at least some of his specimens were collected in proposed as questionable (Boulenger, 1885) and Central Chile (see P. modestus section). Therefore, several insinuated synonymies (Ortiz and Núñez, it cannot be discarded that his L. melanopleurus 1986; Ortiz, 1994; Núñez et al., 2000; Pincheira- specimens may have been collected outside of the Donoso and Núñez, 2005, 2007; Troncoso-Palacios, Atacama Desert in one of the various localities 2014). Even more problematic, the lectotype of L. that Philippi visited in the Valparaíso, Santiago, melanopleurus has four scales on the frontal area, 2 Los Ríos, and Los Lagos regions of Chile (Barros- anterior and 2 posterior, which is a very uncommon Arana, 1904). If this is the case, it would be almost feature in Liolaemus (see Pincheira-Donoso and impossible to determine the taxonomic identity of L. Núñez, 2005), not found in any of the specimens melanopleurus, due to the precarious state of conser- that we reviewed (see Appendix I) except the L. vation of the type specimens, which can explain why melanopleurus lectotype itself. This suggests that the more than 100 years later, no author has been able to lectotype could be an aberrant specimen, as has been

182 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) pointed out for other type specimens of currently match Philippi´s (1860) locality information (Papo- unidentified Liolaemus (Borczyk and Skawinski, so). Ortiz and Núñez (1986) further pointed out that 2019). We conclude that it is not currently possible to NMW 18914: 1,2 are juveniles of L. nigromaculatus. determine to which population of Liolaemus lizards They also proposed that the drawing of P. pallidus the specimens described as L. melanopleurus belong. in Philippi (1860) resembles Liolaemus platei Wer- In regards to the nomenclature, without doubt ner, 1898, but stated that more evidence is needed. L. melanopleurus is an available name because it has However, Ortiz and Núñez (1986) finally considered a description and the types stand. Besides, a nomen P. pallidus a junior synonym of another Philippi spe- dubium is an available name whose application is cies, L. melanopleurus. Recently, Marambio-Alfaro unknown or doubtful, because the type is not iden- and Troncoso-Palacios (2014) agreed with Ortiz tified (ICZN, 1999, Art. 8) or because taxonomic and Núñez (l.c.) that P. pallidus could be related to identity of a species name cannot be determined L. platei, because as of 2014, the only Liolaemus spe- from its existing name-bearing type (ICZN, 1999, cies recorded in Paposo was L. platei (Ortiz, 1973); Art. 75.5). In the case of L. melanopleurus, although however, given that several species related to L. platei type specimens do not allow the taxonomic identi- have been described with type localities geographi- fication of it as junior or senior synonym of one of cally closer to Paposo than the type locality of L. the currently known Chilean Liolaemus species and platei (Donoso-Barros, 1974; Ortiz, 1987;, Núñez et no new specimens have been found, the possibility al., 2001; Marambio-Alfaro and Troncoso-Palacios, of L. melanopleurus as extinct species or aberrant 2014), it seems to be unlikely that the Ortiz`s (1973) specimens cannot be discarded. Liolaemus from Paposo actually belongs to L. platei, being more likely that it belongs to another species of Proctotretus pallidus (Philippi 1860, p. 166, tab. VI, the L. platei group. Moreover, Ruiz de Gamboa and Fig. 3; reproduced here in Fig. 2) Ferrú (2015) subsequently recorded L. nigromacu- Philippi (1860) described and illustrated P. palli- latus from Paposo, so at present two Liolaemus are dus, with the stated type locality of “… bei Paposo”, know from this locality: L. nigromaculatus and one Antofagasta region, Chile, stating that this species Liolaemus sp. of the L. platei group. is abundant but without listed type specimens. It is Photographs of the NMW specimens allow us described as a lizard with a lateral neck fold covered to conclude that the NMW 18914 series consists of by granular scales, with ovate and slightly keeled two species: NMW 18914:1 is a member of the L. dorsal scales; color whitish with mottling of small platei group, whereas NMW 18914:2 is assignable to black spots. Measures are provided in inches (zoll) L. nigromaculatus, which matches with both species and lines (linien), which converted to centimeters currently know from Paposo. In fact, NMW 18914:2 are: total length= 13.4 cm, head length= 1.5 cm, has slightly keeled dorsal scales without mucron, and foreleg length= 2.5 cm, hind leg length= 3.2 cm, tail the nasal scale is separated from the rostral scale as length= 7.6 cm. in L. nigromaculatus (Troncoso-Palacios and Garín, Boulenger (1885, p. 147) considered Procto- 2013). In contrast, NMW 18914:1 shows strongly tretus pallidus a junior synonym of L. nigromacula- keeled dorsal scales with mucron and, although tus. The status of the P. pallidus syntypes has been scales of the snout are a bit damaged, the rostral and disputed. Tiedemann and Häupl (1980) listed two the nasal scales appear to be in contact as occurs in syntypes of P. pallidus (NMW 18914:1,2) with the the species of the L. platei group (Marambio-Alfaro locality of Huasco, Chile, and which were donated and Troncoso-Palacios, 2014). by Franz Steindachner in 1874. We remark that this In regards to the qualitative characters mentio- date is congruent with the 1916 catalogue of the ned by Philippi (1860:166), NMW 18914:1 does not lizards placed in the Chilean National Museum of match those provided for P. pallidus because it has Natural History (Quijada, 1916) in which P. pallidus black dorsal patches (versus mottling of black spots is not mentioned. Tiedemann and Häupl (1980) cite in P. pallidus) and features a black lateral stripe not Peters and Donoso-Barros (1970) as reference for mentioned by Philippi. Moreover, it has strongly the current status of P. pallidus, viz., as synonym of keeled dorsal scales (slightly keeled in P. pallidus) L. nigromaculatus. Ortiz and Núñez (1986) rejected and an inconspicuous lateral neck fold. However, the type status of NMW 18914: 1,2 because the gi- this specimen clearly resembles the drawing of P. ven locality for these specimens (Huasco) does not pallidus provided by Philippi (1860), since both

183 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860) shared a dark lateral stripe, antehumeral spot and collection data of Helocephalus nigriceps, another very similar dark head transversal stripes (Fig. 6). Philippi`s species were similarly confused following However, specimen NMW 18914:2 matches the its publication date (see below). qualitative description of P. pallidus in that it also As occurs with P. melanopleurus, Philippi has small black spots on dorsum, ovate and slightly (1860) based his description of P. pallidus on more keeled dorsal scales, and a lateral neck fold covered than one specimen, but it this case, each specimen is by granular scales. Nevertheless, it does not resemble of a different species. The evidence suggests that he the drawing of P. pallidus provided by Philippi (1860). based the drawing and measurements on specimen In regards to the measurements, NMW 18914:1 NMW 18914:1, but the qualitative characters of dorsal is very close to the data provided by Philippi (1860) scales, dorsal color, and lateral neck folds are based on for P. pallidus: total length= 12.9 cm, head length= NMW 18914:2. We designate the NMW 18914:2 as 1.4 cm and tail length= 7.8 cm. However, NMW lectotype of Procotretus pallidus, because this speci- 18914:2 has very different total length (8.5 cm), head men matches the qualitative characters of the original length (0.9 cm) and tail length (3.5 cm). description and it also the only specimen to present We conclude that P. pallidus is based on both the pale dorsal color used by Philippi to name the NMW 18914 specimens and that the locality infor- species (pallidus means pale). Thus, specimen NMW mation was altered in the NMW collections at some 18914:1, while recognized here as a paralectoype, time following Boulenger’s (1885) synonymy with cannot be a name-bearing type of L. pallidus because L. nigromaculatus and following the restriction of it belongs to unidentified species of theL . platei group. the type locality of L. nigromaculatus to Huasco by Accordingly, P. pallidus becomes a subjective junior Müller and Hellmich (1933a). However, given that synonym of L. nigromaculatus, leaving the current Philippi never went to Huasco (Ortiz and Núñez taxonomy of this species and all valid names for the 1986), we propose to correct the collection locality species of the L. platei group unaffected. of the NMW 18914 specimens to “Paposo, Antofa- gasta Region, Chile”, following Philippi’s (1860:167) Microlophus lessoni Duméril and Bibron, 1837 (Phi- statement that this species was “common at Paposo”. lippi 1860, p. 167) Such alterations of collection data are not unusual, Philippi’s account includes lizards that he assigned given the old dates involved, and we note that the to M. lessoni and notes them as being “common on

Figure 6. Drawing of P. pallidus by Philippi (1860) and photographs of the type specimens. A) Drawing of P. pallidus. B) Specimen NMW 18914:2, here designated as lectotype of P. pallidus and probably used by Philippi (1860) for character description. It is assign- able to L. nigromaculatus. C) Specimen NMW 18914:1 of Liolaemus sp. of the L. platei group, probably used by Philippi (1860) for the drawing of P. pallidus. D) Head of NMW 18914:2. All photographs by Alice Schumacher and Georg Gassner.

184 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) cliffs along the whole coast of the desert”, without February 1855, then the lectotype was collected this further details or specific localities. According to year and most likely, the year was confused with the Mertens (1956), M. lessoni is a junior synonym of year of publication of Philippi`s (1860) book. M. peruvianus (Lesson, 1830), but the latter species does not occur in Chile (Ortiz, 1980; Ruiz de Gam- Aporomera ornata Duméril and Bibron, 1839 (Phi- boa, 2016; Núñez et al., 2018). However, two other lippi 1860, p. 168) Microlophus species occur in the localities visited by Philippi (1860) attributed species authority to Du- Philippi, viz M. maminensis (Donoso-Barros, 1966) méril and Bibron and listed as junior synonyms and M. marianus (Donoso-Barros, 1966) (Núñez Ameiva oculata D’Orbigny and Bibron, 1847, and and Jaksic, 1992; Troncoso-Palacios, 2018). Using Aporomera ocellata Guichenot, 1848 (in Gay, 1848). the key to the Chilean species of the genus Microlo- We note that all these names are currently con- phus provided by Ortiz (1980), we determined three sidered junior synonyms of Callopistes maculatus specimens of Microlophus collected by Philippi and Gravenhorst, 1838. The characterization provided sent to the Zoologisches Museum Berlin to best fit by Philippi (1860) is not detailed, lacking for ex- M. marianus. The specimens were collected in the ample a description of the color pattern. However, Atacama Desert but the exact locality is unknown. he provided a total length of 15 inches (38.1 cm), All specimens have dark throats. While ZMB 5353 a comparatively large size for a Chilean lizard. He A and C are darkened and the dorsal color pattern stated that the specimens that he found showed some is not discernible, ZMB 5353 B (Fig. 7) has dark differences in regards to the Gay (1854) plate, but he paravertebral coloration. These features match M. thinks that these differences are due to the fact that marianus according to Ortiz (1980). Moreover, the painter did not notice certain details. Indeed, M. marianus inhabits in the coast of the Atacama Gay’s (1854) plate of A. ocellata clearly corresponds desert, whereas M. maminensis occurs in the inner to the lizard currently known as C. maculatus, char- Desert (Ortiz, 1980). Thus, we conclude that Philippi acterized by Donoso-Barros (1966: 378) as a large (1860:167) refers to M. marianus. lizard with rounded dorsal black spots bordered by white and brown dorsal color. Moreover, we Helocephalus nigriceps (Philippi 1860, p. 167, tab. VI, reviewed three specimens labeled as Aporomera or- Fig. 1; reproduced here in Fig. 2) nata and collected by Philippi in “Santiago de Chile” Philippi (1860:167) described this lizard from Pajo- (ZMB 5354 [two specimens] and 5355) which can nal, Atacama Desert, Chile. Koslowksy (1898) assig- be unambiguously assigned to C. maculatus (Fig. 9), ned this taxon to Liolaemus signifer var. nigriceps and confirming the species identification. Donoso-Barros (1966: 335) considered it member of Ctenoblepharis. Laurent (1984) definitively returned Final remarks it to Liolaemus; Lobo et al. (2010) placed it in L. mon- tanus group of the subgenus Eulaemus. According Correct taxonomy is a key aspect of biodiversity and to Ortiz and Núñez (1986), one of the syntypes is conservation studies (Mora et al., 2011; Thomsonet lost and the other remains in the Zoologische Staats- al., 2018). However, sometimes taxonomists have to sammlung München, which they designated as the deal with names whose application is uncertain due lectotype (ZSM 38/1930). The lectotype (Fig. 8) and to the lack of or the deterioration of type specimens, the illustration provided by Philippi (1860) (Fig. 2) the lack of or imprecision of type localities, and/or have allowed the unquestionable identification of ambiguous or inaccurate descriptions (e.g. Borczyk this species as the Liolaemus lizard that occurs in the and Skawinski, 2019). The status of several species high Andean areas of the southeastern portion of the described by Philippi (1860) has been controversial Antofagasta Region of Chile and adjacent areas of (Ortiz and Núñez, 1986; Pincheira-Donoso and the Jujuy and Salta Provinces of Argentina (Donoso- Núñez, 2005), but based on the findings of our re- Barros, 1966; Avila et al., 2013) with a black head, search we provide updates on their nomenclatural gray dorsal ground color, and transverse orange and taxonomic status (Table 1). We point out that dorsal bars bordered by dark scales (Donoso-Barros, while Philippi’s (1860) lizard descriptions contain 1966; Pincheira-Donoso and Núñez, 2005). Franzen several omissions and mistakes, these kinds of issues and Glaw (2007, p. 224) stated 1860 as date of collec- were common in the early taxonomy of the Chilean tion of the lectotype, but Philippi went to Pajonal in lizards (see Langstroth. 2011; Troncoso-Palacios,

185 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860)

Figure 7. Specimen ZMB 5353B listed as Microlophus lessoni by Philippi (1860) but here designated as M. marianus. All photo- graphs by Frank Tillack.

186 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021)

Figure 8. Lectotype ZSM 38/1930 of Helocephalus nigriceps (currently Liolaemus nigriceps). Photograph by Michael Franzen.

Figure 9. Specimen ZMB 5355 of Aporomera ornata (currently Callopistes maculatus) collected by Philippi. Photograph by Frank Tillack.

187 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860)

Table 1. Nomenclatural status and taxonomic status of the lizards described by Philippi in the Zoology Section of his book. * There are two specimens with the voucher code MNHNCL 1646. Nomenclatural status Taxonomic status according Type specimens to this and previous research (see text) Helocephalus Lectotype ZSM 38/1930 (extant) and paralectotype Available name Liolaemus nigriceps nigriceps lost (unknown voucher code) Proctotretus Available name but Invalid Lost (unknown voucher code) marmoratus secondary homonym Invalid, junior synonym of Lectotype ZMB 5350 and paralectotypes ZMB P. modestus Available name Liolaemus bellii 70546–47 (all extant) Lectotype MNHNCL 1646* (larger specimen), P. Taxonomic application is Available name paralectotype MNHNCL 1646 (smaller specimen) and melanopleurus unknown paralectotype FMNH 9969 (all extant) Lectotype NMW 18914:2 (extant). NMW 18914:1 Invalid, junior synonym of P. pallidus Available name (extant) cannot be considered a name-bearing type be- Liolaemus nigromaculatus cause it belongs to Liolaemus sp. of the L. platei group.

2018; Troncoso-Palacios et al., 2019). Finally, we re- Literature cited mark that Philippi (1860) did not provide additional Abdala, C.S. & Quinteros, A.S. 2014. Los últimos 30 años de taxonomic information, such as family, for any of estudios de la familia de lagartijas más diversa de Argentina. Actualización taxonómica y sistemática de Liolaemidae. the lizards listed, but the current families are: Fam- Cuadernos de Herpetología 28: 55–82. ily Teiidae, genus Callopistes (listed as Aporomera); Avila, L.J.; Martínez, L.E. & Morando, M. 2013. Checklist Family Tropiduridae, genus Microlophus; and Family of lizards and amphisbaenians of Argentina: an update. Liolaemidae, genus Liolaemus (listed as Helocephalus Zootaxa 3616: 201–238. Barros-Arana, D. 1904. Don Rodolfo Amando Philippi su vida and Proctotretus). i sus obras. Imprenta Cervantes, Santiago. Benavides, E., Baum, R., McClellan, D. & Jack, W. 2007. Acknowledgments Molecular phylogenetics of the lizard genus Microlophus To Mario Penna (Universidad de Chile) for his sup- (Squamata: Tropiduridae): aligning and retrieving indel port. We are grateful to Frank Tillack (Museum für signal from nuclear introns. Systematic Biology 56: 776–797. Bell, T. 1843. . In: Darwin, C. (Ed.). The Zoology of Naturkunde, Berlin, Germany) for sharing data, the Voyage of the H.M.S. Beagle, Under the Command photographs of several specimens collected by of Captain Fitzroy, R.N., During the Years 1832 to 1836. Philippi, literature and for his very appreciate cor- Volume 5: 1–51. London: Smith, Elder and Co. rections and comments to the early version of this Borczyk, B. & Skawinski, T. 2019. Tracking down the lizards from Gravenhorst’s collection at the University of Wroc1aw: manuscript. Herman Núñez (Museo Nacional de type specimens of Callopistes maculatus Gravenhorst, 1838 Historia Natural de Chile, Santiago), Jorge Artigas and three Liolaemus species rediscovered. PeerJ 7:e6525. (Museo de Zoología de la Universidad de Concep- Boulenger, G.A. 1885. Catalogue of the lizards in the British ción, Chile), Patricio Zavala (Colección Patricio Museum (Natural History). Volume 2. , Xenosauriae, Sánchez de la Pontificia Universidad Católica de Zonuridae, Anguidae, Annellidae, Helodermatidae, Varanidae, Xantusiidae, Teiidae. Printed by Order of the Chile, Santiago) and Franklin Troncoso (Museo Trustees, London. de Historia Natural de Concepción) for access to Donoso-Barros, R. 1961. The reptiles of the Lund University specimens under their care. Kathleen Kelly (Field Chile Expedition. Copeia 1961: 486–487. Museum of Natural History, Chicago, USA), Michael Donoso-Barros, R. 1966. Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago. Franzen (Zoologische Staatssammlung München, Donoso-Barros, R. 1974. Nuevos reptiles y anfibios de Chile. Germany), Alice Schumacher and Georg Gassner Boletín de la Sociedad de Biología de Concepción 48: 217–229. (Naturhistorisches Museum Wien, Austria) kindly Etheridge, R.E. 1995. Redescription of Ctenoblepharys adspersa send us photographs of type specimens collected by Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: R.A. Philippi. One anonymous reviewer, Roberto Squamata: Tropiduridae). American Museum Novitates 3142: 1–34. Langstroth and the editor Sebastian Quinteros for Franzen, M. & Glaw, F. 2007. Type catalogue of reptiles in the all their corrections and comments. Zoologische Staatssammlung München. Spixiana 30: 201–274.

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Beiträge zur Liolaemus isabelae Navarro and Núñez, 1993 and Liolaemus Zoologie gesammelt auf einer Reise um die Erde. Siebente nigroventrolateralis Ortiz 1994 (Iguania: Tropiduridae: Abhandlung. Amphibien. Nova Acta Physico-Medica Liolaeminae) from northern Chile. Herpetological Journal Academia Caesarea Leopoldino-Carolina, Halle 17: 185–268. 17: 65–67. Young-Downey, A. & Moreno, J. 1992. A new species of Pincheira-Donoso, D.; Scolaro, J.A. & Lura, P. 2008. A tropidurine lizard (Squamata: Tropiduridae) from Los monographic catalogue on the systematics and phylogeny Andes of northern Chile. Gayana (Zoología) 55: 391–396. of the South American iguanian lizard family Liolaemidae (Squamata, Iguania). Zootaxa 1800: 1–85. Appendix I. Quijada, B. 1916. Catálogo sistemático de los reptiles chilenos i Specimens examined. Abbreviation used: FMNH (Field Mu- estranjeros conservados en el Museo Nacional de Historia seum of Natural History, Chicago, USA), MNHNCL (Museo Natural. Boletín del Museo Nacional de Historia Natural de Nacional de Historia Natural de Chile, Santiago), MRC Chile 9: 22–47. (Museo Regional de Historia Natural de Concepción, Con- Ruiz de Gamboa, M. 2016. Lista actualizada de los reptiles de cepción), MZUC (Museo de Zoología de la Universidad de Chile. Boletín Chileno de Her­petología 3: 7–12. Concepción, Concepción), NMW (Naturhistorisches Museum Ruiz de Gamboa, M. & Ferrú, M. 2015. Liolaemus nigromaculatus Wien, Austria), SSUC (Colección de la Pontificia Universidad (Many-spotted Tree Iguana). Herpetological Review 46: 217. Católica de Chile, Santiago), ZMB (Museum für Naturkunde, Szederjesi, T.; Latif, R.; MÁrton, O. & Csuzdi, C. 2018. Berlin, Germany; formerly Zoologisches Museum Berlin), Resurrection of the earthworm species Dendrobaena ZSM (Zoologische Staatssammlung München, Germany) fedtschenkoi (Michaelsen, 1900), a former synonym of Dendrobaena byblica (Rosa, 1893) (Clitellata: Megadrili). Type specimens collected by Philippi: Liolaemus melano- Zootaxa 4496: 190–196. doi: 10.11646/zootaxa.4496.1.14. pleurus. MNHNCL 1646 (2 specimens). Chile. FMNH 9969. Thomson, S.A.; Pyle, R.L.: Ahyong, S.T.: Alonso-Zarazaga, M.; Chile. Liolaemus modestus. ZBM 5350, 70546–47. Santiago Ammirati, J.; et al. 2018. Taxonomy based on science is de Chile (highlands), Metropolitan Region, Chile. Liolaemus necessary for global conservation. PLoS Biology 16: e2005075. nigromaculatus (L. bisignatus holotype). MNHN-CL 1477. Tiedemann, F. & Häupl, M. 1980. Typenkatalog der Atacama (restricted to Caldera, Atacama Region), Chile. herpetologishen Sammlung Teil II: Reptilia. Kataloge Liolaemus pallidus. NMW 18914:1,2. Paposo (previously der wissenschaftlichen Sammlung des Naturhistorischen listed from Huasco in error), Antofagasta Region, Chile. L. Museums in Wien. 4 (Vertebrata 2): 5–70. nigriceps. ZSM 38/1930. Pajonal, Antofagasta Region, Chile. Tschudi, J.J. 1845. Reptilium conspectus quae in Republica Peruana reperiuntur et pleraquae observata vel collecta sunt Additional material (no types) collected by Philippi: Cal- in itinere a Dr. J. J. de Tschudi. Archiv für Naturgeschichte lopistes maculatus. ZMB 5354 (two specimens), 5355. San- 11: 150–170. tiago de Chile, Metropolitan Region, Chile. Microlophus Troncoso-Palacios, J. 2013. Revisión del estatus taxonómico de marianus. ZMB 5353 (A, B and C). Atacama Desert, Chile. Liolaemus donosoi Ortiz, 1975 (Iguania: Liolaemidae). Boletín Specimens not collected by Philippi but from localities visited del Museo Nacional de Historia Natural de Chile 62: 119–127. by Philippi or from the surroundings: Liolaemus atacamensis. Troncoso-Palacios, J. 2014. Nueva lista actualizada de los reptiles MZUC 30193, 30196. Punta Teatinos, Coquimbo Region, terrestres de la Región de Atacama, Chile. Boletín Chileno Chile. J.C. Ortiz coll. September 14, 1982. SSUC Re 454, de Herpetología 1: 1–4 464–68. Lomas de Buitre, Freirina, Atacama Region, Chile. Troncoso-Palacios, J. 2018. Propuesta de restricción de la J. Troncoso-Palacios, Y. Marambio and D. Hiriart colls. May, localidad tipo de Microlophus tarapacensis (Donoso-Barros, 2012. SSUC Re 455–61. Playa Humedal Pachingo, Coquimbo 1966) (Squamata: Tropiduridae) en base a evidencias Region, Chile. C. Garín coll. December 10, 2009. SSUC Re históricas. Cuadernos de Herpetología 32: 123–127. 469. 20 km N from Vallenar, Atacama Region, Chile. F. Ferri Troncoso-Palacios, J. & Garín, C.F. 2013. On the identity of Liolaemus coll. 2010. SSUC Re 470–71. El Trapiche, Coquimbo Region, nigromaculatus Wiegmann, 1834 (Iguania, Liolaemidae) and Chile. J. Troncoso-Palacios, Y. Marambio and D. Hiriart colls.

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May, 2012. Liolaemus bellii. MNHNCL 1599. Sewell, O`Higgins 462–63. 50 km N from Vallenar, Atacama Region, Chile. J. Region, Chile. M. Elgueta coll. December 1982. SSUC 201–05. Troncoso-Palacios coll. June 5, 2011. SSUC Re 476–77. Caldera, Casa de Piedra, Farellones, Metropolitan Re­gion, Chile. F. Ferri Atacama Region, Chile. Y. Marambio coll. May, 2012. SSUC Re coll. October 12, 2010. SSUC Re 206–09. El Colorado, Farel- 478 20 Km SE from Puerto Viejo, Atacama Region, Chile. J. lones, Metropolitan Region, Chile. F. Ferri coll. November Troncoso-Palacios and Y. Marambio colls. May, 2012. SSUC Re 13, 2011. SSUC Re 398–404, 543. El Olivares, Metropolitan 642–43. Paposo, Antofagasta Region, Chile. M. Ferru and M. Region, Chile. C. Garín coll. Undated. SSUC Re 562–66. La Ruiz de Gamboa colls. June, 2014. ZMB 613. Transect between Parva, Metropolitan Region, Chile. J. Opazo coll. December, Puerto Viejo and Copiapó, Atacama Region, Chile. F.J.F. Meyen 2003. SSUC Re 654, 656. Lagunillas, Metropolitan Region, coll. March, 1831. Liolaemus nitidus. SSUC Re 59, 69, 72–74, Chile. D. Esquerré coll. February 15, 2015. Liolaemus constan- 78–79, 82. Altos de Cantillana, Metropolitan Region,­ Chile. zae. MNHNCL 1499–1500. Que­brada de Taltal, Agua Verde, Unknown collector and date. SSUC Re 298. Dunas de Ritoqui, Antofagasta Region, Chile. S. Zunino and M. Riv­eros colls. Valparaíso Region, Chile. Ferri F. coll. November 13, 2010. February 10, 1975. MNHNCL 1516–1520. Quebrada de Taltal, SSUC Re 299, 300. Road to Farellones, curve 20, Metropolitan Agua Verde, Antofagasta Region, Chile. H. Núñez, J. Yáñez and Re­gion, Chile. Ferri F. coll. December 8, 2010. SSUC Re 301. Contreras colls. September 27, 1982. MZUC 29247, 29250–51. Lo Valdés, Región Metropolitana, Chile. Ferri F. coll. January Toconao, Antofagasta Region, Chile. Unknown collector and 9, 2011. SSUC Re 302–03. Road to Farellones, curve 20, Met- date. MZUC 28763–65, 28767–69. Agua Verde (Que­brada de ropolitan Re­gion, Chile. Ferri F. coll. March 15, 2012. SSUC Re Taltal), Antofagasta Region, Chile. J.C. Ortiz, S. Zunino and M. 418. Altos de Cantillana, Metropolitan Re­gion, Chile. C. Correa Riveros colls. February 10, 1975. SSUC Re 338– 39, 341–42, 346. coll. December 14, 2011. SSUC Re 548. Parque Nacional Llanos Cuesta Barros Arana, Antofagasta Region, Chile. F. Ferri coll. de Challe, Atacama Region, Chile. G. Lobos. June, 2002. SSUC October 22, 2011. SSUC Re 340, 343–45, 347. Cuesta Barros Re 705–06. Quebrada de la Plata, Maipú, Metropolitan Region,­ Arana, Antofagasta Region, Chile. F. Ferri coll. October 25, Chile. C. Garín coll. December 17, 2015. SSUC Re 725, 727–28. 2011. SSUC Re 348. Southern Salar de Atacama, Antofagasta Cerro Provincia, Metropolitan Region,­ Chile. C. Garín coll. Re­gion, Chile. F. Ferri coll. October 24, 2011. SSUC RE 482. El Liolaemus paulinae. SSUC Re 361. Calama, Antofagasta Region, Abra, Antofagasta Region, Chile. G. Lobos and F. Torres colls. Chile. C. Garín coll. October 27, 2010. SSUC Re 486. Mina El November 23, 2003. SSUC RE 483, 485, 488. El Abra, Antofa- Abra, Antofagasta Region, Chile. G. Lobos and F. Torres colls. gasta Region, Chile. G. Lobos and F. Torres colls. November 21, November 22, 2003. SSUC Re 487. Mina El Abra, Antofagasta 2003. Liolaemus hellmichi. MNHNCL 4126–37. Cerro Moreno, Region, Chile. G. Lobos and F. Torres colls. November 21, 2003. Antofagasta Region, Chile. P. Espejo coll. December 31, 2000. SSUC Re 561. Embalse rio Loa, Antofagasta Region, Chile. G. MZUC 25942–52. Cerro Moreno, Antofagasta Region, Chile. Lobos and F. Torres colls. November 22, 2003. Liolaemus platei. J.C. Ortiz coll. April 2, 2001. Liolaemus isabelae. SSUC Re 157, MZUC 2152–53. Combarbalá, Coquimbo Region, Chile. I. 159, 160. El Cerrito, Salar de Pedernales, Atacama­ Region, Peña coll. November, 1961. SSUC Re 321. Illapel, near Reserva Chile. F. Ferri and J. Troncoso-Palacios colls. February 22, 2012. Nacional Las Chinchillas, Coquimbo Region, Chile. F. Ferri SSUC Re 158. Montandón, Salar de Pedernales, Atacama Re- coll. Undated. SSUC Re 420. Coquimbo, Coquimbo Region, gion, Chile. F. Ferri and J. Troncoso-Palaci­os colls. February 22, Chile. J. Troncoso-Palacios and Y. Marambio colls. December 2012. Liolaemus juanortizi. MZUC 11782. Río Patón, Atacama 12, 2011. SSUC Re 526, 555. Coquimbo, Coquimbo Region, Region, Chile. T. Cekalovic coll. December 20, 1963. MZUC Chile. J. Troncoso-Palacios and Y. Marambio colls. May, 2012. 11770. Puquios, Atacama (road through Puquios?), Chile. R. Liolaemus velosoi. MZUC 36612–14, 36618–20, 36624. Estación Buzeta coll. December 19, 1963. SSUC Re 755–56. Paso pircas Paipote, Atacama Region, Chile. J.C. Ortiz coll. February 16, coloradas, Copiapó, Atacama Region, Chile. Unknown collector 1978. MZUC 32695, 32699, 32702, 32704, 32706. Copiapó, and date. Liolaemus nigrocoeruleus. SSUC Re 527–32, 552–54. Atacama Region, Chile. R. Moreno coll. February, 2000. MRC Seven km NE from Barranquilla, unamed hill, Atacama Region, 054. Copiapó, Atacama Region, Chile. Unknown collector. Chile. J. Troncoso-Palacios and Y. Marambio-Alfaro colls. May, June 16, 1982. MRC 055. Copiapó, Atacama Region, Chile. 2012. SSUC Re 533–34. Nine km NE from Barranquilla, un- Unknown collector. April 20, 1982. MRC 061, 062, 066. Copi- amed hill, Atacama Region, Chile. J. Troncoso-Palacios and Y. apó. Unknown collector. July 19, 1982. SSUC Re 322–26. Tierra Marambio-Alfaro colls. May, 2012. Liolaemus nigromaculatus. Amarilla. J. Troncoso-Palacios and F. Ferri. colls. November 23, MNHNCL 2237–38. 20 km of Caldera, between Copiapó and 2011. SSUC Re 330. Diego de Almagro. F. Ferri, J. Troncoso- Caldera, Atacama Region, Chile. H. Núñez coll. September Palacios colls. December 9, 2011. SSUC Re 327–29, 331–34, 30, 1991. MNHNCL 2249–55. Travesía, Copiapó, Atacama 419. Diego de Almagro. F. Ferri and J. Troncoso-Palacios colls. Region, Chile. H. Núñez coll. September 28, 1999. MRC 051, December 12, 2011. Liolaemus zapallarensis. MZUC 29118, 053. Caldera, Atacama Region, Chile. J. Moreno coll. May, 29122–23. Las Tacas, Coquimbo Region, Chile. J.C. Ortiz coll. 1982. MRC 162, 273, 276, 282–83. Caldera, Atacama Region, October 15, 1976. MZUC 29127. Las Tacas, Lagunillas, Coqui- Chile. J. Moreno coll. 1983. MRC 087–94. Copiapó, Atacama mbo Region, Chile. J.C. Ortiz and J. Simonetti colls. September Region, Chile. C. Valdovinos coll. September 15, 1984. MRC 10, 1977. SSUC Re 472. Totoralillo, Coquimbo Region, Chile. J. 514. Copiapó, Atacama Region, Chile. Unknown collector. Troncoso-Palacios and Y. Marambio colls. December 12, 2011. November, 1996. MZUC 14820. Sector algarrobal (between Vallenar and Copiapó), Atacama Region, Chile. J. Moreno coll. © 2021 por los autores, licencia otorgada a la Asociación Herpetológica Argentina. Este artículo es de acceso abierto y distribuido bajo los términos y condiciones de August 21, 1984. SSUC Re 306–15, 474–75. Caldera, Atacama una licencia Atribución-No Comercial 2.5 Argentina de Creative Commons. Para ver Region, Chile. F. Ferri coll. November, 2011. SSUC Re 453, una copia de esta licencia, visite http://creativecommons.org/licenses/by-nc/2.5/ar/

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